An Integrated Multilayer Network Orchestrating Fruit Softening: From Hormonal Crosstalk to Epigenetic Regulation

Fruit softening determines postharvest quality but also drives postharvest losses. Despite extensive research, a fully integrated view of its underlying molecular network remains lacking. Drawing on studies across climacteric and non‑climacteric fruits, we present a multi‑tiered regulatory framework encompassing hormonal signaling, transcriptional hierarchy, epigenetic modulation, and cooperative cell wall disassembly. We synthesize current knowledge into a cohesive model that emphasizes crosstalk and feedback across these layers, critically assess persistent gaps including limited cross‑species conservation and unresolved cellular heterogeneity, and outline how emerging technologies such as single‑cell multi‑omics, three‑dimensional genomics, and precision genome editing can transform our ability to predict and engineer fruit texture. A key paradigm emerging from this synthesis is the reciprocal regulation between transcription factors and the epigenetic landscape, which stabilizes the ripening state and serves as a critical interface for environmental signal integration. This integrated framework identifies key regulatory nodes for targeted engineering and provides a roadmap for designing cultivars and postharvest strategies that minimize losses while enhancing fruit quality.